Effect of Mating Disruption on Reproductive Behavior in the European Pine Sawfly, Neodiprion sertifer (Hymenoptera: Diprionidae)

Females of the European pine sawfly, Neodiprion sertifer (Geoffr.) (Hymenoptera: Diprionidae), were released and observed inside and outside (=control) of areas treated with their sex pheromone for the purpose of mating disruption. In 1992 and 1993, respectively, 0 and 2.5% of the females were observed mating in the pheromone treated area during the first day, compared with 28 and 26% in the control area. Of the females that mated, approximately 50% stayed on their twigs and oviposited, while the remaining 50% disappeared. Significantly more females disappeared from their twigs inside the treated area compared to the control area in both years. No difference was found in the proportion of unmated females ovipositing (2–12%) between the treated and the control area. Predation was responsible for much of the disappearance, but deliberate dispersal was also noted. Mating and dispersal in female N. sertifer are discussed in relation to population density and overall fitness.     

Geographic variation in winter freezing susceptibility in the eggs of the European pine sawfly (Neodiprion sertifer)

Abstract 1 The European pine sawfly, Neodiprion sertifer (Geoffroy) (Hymenoptera, Diprionidae), frequently defoliates Scots pine (Pinus sylvestris L.) forests in northern Europe. It overwinters as an egg. It has been proposed that the high egg mortality caused by low winter temperatures limits the occurrence of outbreaks to the southern part of Fennoscandia. 2 In this study, variation in freezing avoidance by egg supercooling between four Finnish populations (originating between latitudes 60°N and 69°N) of N. sertifer was tested by differential thermal analysis. Offspring of 20 females within each population were selected for the study. The freezing avoidance of parasitized eggs was also examined. 3 The northernmost Inari population was found to be the cold hardiest, and the southernmost (Hanko) was the least hardy population. The within‐population variation between females was greatest in the population from Inari, and the next greatest in the one from Hanko. The inland populations in Eastern Finland had the smallest within‐population variation in freezing avoidance. 4 The high variation in freezing avoidance of eggs will enable N. sertifer to adapt to the predicted climate change and to spread its distribution northwards. This may also change the risk for outbreaks in this area. Parasitized eggs froze at higher temperature than healthy eggs. This observation indicates that N. sertifer may experience reduced egg parasitism in certain winter climate conditions.     

Monitoring the European pine sawfly with pheromone traps in maturing Scots pine stands

Abstract

• 1 During 1989–93, field studies were conducted in Finland to develop a method based on pheromone traps to monitor and forecast population levels of the European pine sawfly (Neodiprion sertifer Geoffr.) and tree defoliation.
• 2 Three traps per site were baited with 100 µg of the N. sertifer sex pheromone, the acetate ester of (2S,3S,7S)‐3,7‐dimethyl‐2‐pentadecanol (diprionol), in maturing pine stands in southern and central Finland. In addition, three different dosages (1, 10 and 100 µg) of the pheromone were tested in 1991–92.
• 3 The highest number of males was observed in traps baited with the highest dose. On average, there was a 10‐fold increase in trap catch between lure doses.
• 4 Density of overwintering eggs was used to evaluate the effectiveness of pheromone traps in predicting sawfly populations. The proportion of healthy overwintering eggs was determined each year. A model based on the number of current shoots on sample trees, diameter at breast height and tree height was formulated to estimate eggs per hectare.
• 5 Linear regression analysis produced high coefficients of determination between number of males in traps and density of total eggs in the subsequent generation, when populations were at peak densities. The relationships were not significant for low population densities. The results indicate a risk of moderate defoliation when the seasonal trap catch is 800–1000 males per trap or higher.

     

Sex pheromone perception in male pine sawflies,Neodiprion sertifer (Hymenoptera; Diprionidae)

Summary Electroantennographic and single sensillum recordings were performed on male pine sawfly, Neodiprion sertifer, antennae. Responses to the sex pheromone component (2S, 3S, 7S)- 3,7-dimethyl-2-pentadecenyl (diprionyl) acetate (SSS:OAc), to the behavioral inhibitor (2S, 3R, 7R)-diprionyl acetate (SRR:OAc), to the six other enantiomers of diprionyl acetate, and to the biosynthetic precursor diprionol were recorded. Responses to trans-perillenal, a monoterpene identified in female gland extracts and to (2S, 3S, 7S)-diprionyl propionate (SSS:OPr), a field attractant for N. sertifer and some related sawfly species were also recorded.EAG recordings demonstrated a high antennal sensitivity to SSS:OAc and to SSS:OPr. A somewhat lower response was elicited by SRR:OAc.Single sensillum recordings revealed 8–12 different cells firing in each sensillum, corresponding to the number of cells observed in earlier morphological investigations. Out of these cells all, except one, responded to SSS:OAc and to SSS:OPr. No differences in the response to the two components could be observed. The largest amplitude cell in each sensillum was specifically tuned to the behavioral antagonist, SRR:OAc. The pheromone perception system encountered in male pine sawflies thus differs clearly from that observed in moths.Abbreviation EAG electroantennogram - OAc acetate - OPr propionate     

Interspecific competition between the principal larval parasitoids of the pine sawfly,Neodiprion sertifer (Geoff.) (Hym.: Diprionidae)

Summary Interspecific competition between the 3 principal larval parasitoids of the pine sawfly, Neodiprion sertifer, is of common occurrence when total larval parasitism, and hence multiple attack rates, are high. At the intrinsic level, the ectoparasitoid Exenterus abruptorius is superior to the 2 endoparasitoids, Lophyroplectus luteator and Lamachus eques, respectively, whereas L. luteator is superior to L. eques only. During mass outbreaks of the host competition between Exenterus and Lophyroplectus is most intense, whilst Lamachus fails to build up in the presence of its 2 competitors (Fig. 1). However, due to its superior host finding ability, Lamachus dominates in areas of low host densities where the 2 other species are less efficient (Table 1). L. luteator frequently sustains heavy losses in direct competition with E. abruptorius, but these can be largely compensated in the next generation because its fertility is about 5-times higher than that of Exenterus.     

Fecundity of the European pine sawfly Neodiprion sertifer (Hymenoptera, Diprionidae) related to cyclic outbreaks: Invariance effects

At the end of ontogenesis, individual variability of the fecundity of females of Neodiprion sertifer constitutes 30–120 eggs (with rare fluctuations from 16 to 140 eggs). Average fecundity as an adaptive index in a certain biocenosis, however, depends on many factors, including the structural organization of the sawfly population. The presence of a long diapause in some individuals results in the cleavage of the offspring into groups with different dates of reactivation. At the same time, mainly heavy eonymphs, giving rise to large females, stay in the diapause. Their participation in the reproduction together with females, raised in other years under different conditions, equalize the dimensional structure of the breeding fraction of the population. During population waves, revealed during 35 years of my continuous observations, fecundity fluctuated closely to the average value with deviations constituting 19–25%. It hardly depends on the population density of the phyllophagan. The intraspecific structuredness of the population in relation to the size of females and its transformation by the moment of their emergence promotes the evolutionary stability of the degree of fecundity of the sawfly.     

Development of mating disruption for control of pine sawfly populations

Mating disruption of the pine sawflyNeodiprion sertifer (Geoffroy) (Hymenoptera: Diprionidae) was strongly indicated by reduced male trap catches in pine plantations permeated with the sex phermone, (2S, 3S, 7S)-diprionly acetate. The trap catch reduction was 95 to near 100% when dispensers every 10 m were used, giving a total release of about 3 mg per hectare and day. Two mg of pheromone per cotton roll dispenser maintained low catches for the whole season (about 2 months) without any renewal of disruption dispensers. Anerythro-mixture was as effective as the pure pheromone isomer. The effects of the experiments on population density and sex ratio were not possible to investigate, due to a general collapse of the population, also outside experimental plots, the year after the experiments.     

Field response of the pine sawfly Neodiprion sertifer to the pheromone (2S,3S,7S)-diprionyl acetate and its stereoisomers

All eight optical isomers of 3,7-dimethyl-2-pentadecanyl acetate (diprionyl acetate), of high optical purity (> 97.4%), were tested for a behavioural activity on male pine sawflies, Neodiprion sertifer (Geoffr.) (Hymenoptera: Diprionidae), in northern Europe. Males were strongly attracted to (2S,3S,7S)-diprionyl acetate. Addition of more than 0.1% of the (2S,3R,7R)-isomer reduced the catch and above 2% the attraction was completely inhibited. Contrary to what has been reported for North American and Japanese populations, so significant synergistic effect of small amounts of the (2S,3R,7R)-isomer could be demonstrated. The effects of addition of the other six optical isomers alone or in combinations, were also studied, but none was found to be a synergist. The (2S,3R,7S)-isomer had a weak inhibitory effect, and completely inhibited the attraction to the (2S,3S,7S)-isomer when applied in about equal amounts as the attractant. In some cases a reduction in catch was noted when other isomers were tested, but this could be attributed to the very small amounts of the inhibitory (2S,3R,7R)-isomer present in these isomers.     

中国吉松叶蜂属一新种记述（膜翅目：松叶蜂科）

记述采自陕西秦岭的吉松叶蜂属Gilpinia Benson1新种－青扦吉松叶蜂G.wilsonae sp.n。该种昆虫青扦Picea wilsonii Mast.，一年发生两代，结茧过冬。新种的模式标本保存于西北林学院昆虫标本室。     

Geographic variation in the field response of male European pine sawflies, Neodiprion sertifer, to different pheromone stereoisomers and esters

The European pine sawfly, Neodiprion sertifer (Geoffroy) (Hymenoptera: Diprionidae), is a widespread and economically important forest insect. The sex pheromone communication system of this species has been previously investigated in North America, Japan and Europe, with the acetate or propionate of the alcohol (2S,3S,7S)-3,7-dimethyl-2-pentadecanol (diprionol) shown to be the main pheromone component. In some locations, male attraction either increased or decreased by the addition of the (2S,3R,7R)-diprionyl acetate isomer. However, these studies were made with different batches of synthetic pheromones, with different types of traps and according to different procedures, so the observed differences might not reflect true geographic variation. Here we investigate the geographic pattern of male sawfly response by using identical chemicals, traps and experimental procedures at eight field sites ranging from Japan in the east to Canada in the west. We found an increased inhibitory effect of the (2S,3R,7R)-isomer from Japan and Siberia to Europe. At the eastern sites, increasing amounts of the (2S,3R,7R)-isomer up to and equal to the amount of the (2S,3S,7S )-isomer, did not influence the trap catch, whereas at sites in Europe, as little as 1% of the (2S,3R,7R)-isomer almost completely inhibited the attraction. The response of the North American population was intermediate. The only site in which the (2S,3R,7R)-isomer was essential for the attraction of males was in Siberia. A similar pattern was found for the (2S,3R,7S)-isomer. Both the acetate and the propionate form of the (2S,3S,7S)-isomer were attractive by themselves in Japan, Europe and North America, and neither the (2S,3R,7S)-isomer nor the (2S,3R,7R)-isomer alone were attractive, in the acetate or propionate form. We discuss the significance of our findings for the development of more efficient monitoring schemes and for the causes of population divergence and speciation in the European pine sawfly.     

Long-term effects of exogenous methyl jasmonate application on Scots pine (Pinus sylvestris) needle chemical defence and diprionid sawfly performance

Scots pine [ Pinus sylvestris L. (Pinaceae)] trees with four different seed origins were exposed to exogenous applications of the elicitor, methyl jasmonate (MeJA), for three consecutive years. We studied the effects of MeJA on needle chemistry (including monoterpenes, sesquiterpenes, and tricyclic resin acids), plant growth, and the performance of two diprionid sawflies, the European pine sawfly ( Neodiprion sertifer Geoffr.) and the common pine sawfly ( Diprion pini L.) (both Hymenoptera: Diprionidae). In general, foliar MeJA application affected the whole range of needle secondary chemistry with significantly higher concentrations of two monoterpenes, β-pinene and limonene, in particular. Furthermore, for some seed origins the growth rates of N. sertifer and D. pini larvae were lower on needles of MeJA-treated plants with either high total terpene or high resin acid concentrations. However, inconsistencies in diprionid sawfly performance within each studied Scots pine origin suggest genetic variance in needle secondary chemistry. The differences between selected seed origins and notably variable responses to MeJA application imply that adaptation of the seed to new conditions may have had an impact on secondary chemistry and, thus, on insect performance. Finally, our results suggest that modification of Scots pine defence by a low-concentration exogenous elicitor affects the production of terpenoids in the newly growing needles, leading to poorer pine sawfly performance in origins with high terpenoid content, while not harming the growth of Scots pine trees.     

Mitonuclear discordance is caused by rampant mitochondrial introgression in Neodiprion (Hymenoptera: Diprionidae) sawflies

We investigate the pervasiveness of hybridization and mitochondrial introgression in Neodiprion Rohwer (Hymenoptera; Diprionidae), a Holarctic genus of conifer-feeding sawflies. A phylogenetic analysis of the lecontei species group revealed extensive discordance between a contiguous mitochondrial region spanning three genes (COI, tRNA-leucine, and COII) and three nuclear loci (EF1alpha, CAD, and an anonymous nuclear locus). Bayesian tests of monophyly and Shimodaira-Hasegawa (SH) tests of topological congruence were consistent with mitochondrial introgression; however, these patterns could also be explained by lineage sorting (i.e., deep coalescence). Therefore, to explicitly test the mitochondrial introgression hypothesis, we used a novel application of coalescent-based isolation with migration (IM) models to measure interspecific gene flow at each locus. In support of our hypothesis, mitochondrial gene flow was consistently higher than nuclear gene flow across 120 pairwise species comparisons (P < 1 x 10(-12)). We combine phylogenetic and coalescent evidence to identify likely cases of recent and ancient introgression in Neodiprion, and based on these observations, we hypothesize that shared hosts and/or pheromones facilitate hybridization, whereas disparate abundances between hybridizing species promote mitochondrial introgression. Our results carry implications for phylogenetic analysis, and we advocate the separation of high and low gene flow regions to inform analyses of hybridization and speciational history, respectively.     

Fecundity and egg viability in relation to female body size in Neodiprion sertifer (Hymenoptera: Diprionidae)

Relationships between female size and fecundity, egg viability and embryonal development were studied in Neoriprion sertifer. Sawfly females, reared from the 1st instar larvae on Scots pine needles heavily affected by industrial pollutants or on relatively unpolluted needles, were induced to lay eggs parthenogenetically in the field in Finland. Females reared on polluted needles were smaller and oviposited an average of 68.8 eggs, and those reared on unpolluted needles 79.4 eggs per female. However, the higher egg viability in the former group masked the effect of reduced female size. This compensation was primarily related to the size of the females, larger individuals producing relatively fewer viable eggs in both groups.     

Infection of a nucleopolyhedrovirus to Neodiprion zhejiangenis Zhou & Xiao (Hymenoptera: Diprionidae)

Neodiprion zhejiangenis Zhou &; Xiao 1981 (Hymenoptera: Diprionidae) nuclear polyhedrosis virus (NezhNPV) is a biocontrol factor with high development potential and application prospects. To study the insecticidal activity of NezhNPV against the larvae of N. zhejiangenis, the susceptibility of larvae at different instars to different NezhNPV concentrations was determined based on laboratory bioactivity. At 2.0?×?10⁷ polyhedron inclusion body (PIB)/mL, the median lethal time required to kill 50% of the instar 2 and 3 larvae was approximately 5 d, whereas the median lethal time of the instar 4 and 5 larvae was 8–9 d. Interestingly, at the NezhNPV concentrations of 2.0?×?10³ and 2.0?×?10⁴ PIB/mL, the feeding ability of the surviving larvae was decreased by 57.4% and 76.4%, respectively, compared with the controls; the pupal weight at both concentrations was decreased by approximately 27%, and the adult emergence rate was decreased by 27.4% and 50.9%, respectively, with a significantly higher proportion of males than of females. The results showed that younger instar larvae of N. zhejiangenis were more susceptible to NezhNPV infection than older instar larvae. The larval mortality rate was dependent on the larval instar and NezhNPV concentration.     

Mating disruption of pea moth (Cydia nigricana) in organic peas (Pisum sativum)

We appraised mating disruption (MD) to control pea moth, Cydia nigricana (Fabricius) (Lepidoptera: Tortricidae), by assessing male attraction to monitor traps, larval pod infestation, and larval age structure in pheromone‐treated and untreated grain pea fields [Pisum sativum L. (Fabaceae)], over a 5‐year period. Cellulose pheromone dispensers were manually attached to the top shoots of pea plants and released 540 mg ha^?1 day^?1 synthetic pheromone E8,E10‐dodecadien‐1‐yl acetate in a first test series (2000–2001) and ca. 4 200 mg pheromone ha^?1 day^?1 in a second series (2004–2006). The dispensers had a half‐life of about 30 days. Although male attraction to pheromone monitoring traps was largely suppressed at the edges and within MD fields in both test series, MD treatments did not reduce pod infestation in the open field in 2000 and 2001. In the 2004–2006 series, larval damage reduction was achieved in the majority of the trials but overall MD efficacy in the open field was only 61% and not significant. In contrast, in field cages placed within the experimental sites and supplied with unmated pea moths, MD control was consistently high and significant. There were no obvious differences in the larval age distribution in all MD and control treatments, suggesting that infestations started and developed further similarly. As a univoltine species, C. nigricana larvae stay in the soil of pea fields for hibernation and pupate. The following year, emerging adults disperse and fly to the closest pea crop. Combined emergence site and pea crop treatments were conducted over 2 years to include this early migration phase of C. nigricana adults. However, the emergence site treatments did not enhance MD‐control efficacy. We conclude that mating activity was only prevented in cage tests, whereas substantial mating occurred during the transit phase outside the pheromone‐treated fields either within non‐crop vegetation and/or at the edges of pheromone‐treated pea fields orientated upwind. Thus, resulting gravid female entry can be regarded as the major constraint to reliable MD control.     

Influence of host plant vs. natural enemies on the spatial distribution of a pine sawfly, Neodiprion autumnalis

Abstract .1. The pine sawfly, Neodiprion autumnalis , infests ponderosa pine, Pinus ponderosa , growing at low densities near the bottom of an altitudinal gradient in Arizona, U.S.A. The relative importance of host-plant quality vs. natural-enemy effects in determining the spatial distribution of this sawfly was examined over a 3-year period.
2. Field and laboratory bioassays were conducted on all life stages of N. autumnalis at two forest stand densities (high ≥ 23 m² ha^–1, low ≤ 7 m² ha^–1) and at two elevations (bottom slope = 2390 m, top slope = 2540 m). These experiments were used for constructing life tables of N. autumnalis that compared the effects of host-plant quality on oviposition preference and progeny performance with the effects of natural enemies at different tree densities and elevations.
3. Life-table analyses determined that mortality attributed to host-plant effects during the egg and larval stages had the largest impact on fitness between tree densities and elevations.
4. Natural enemies caused a significant reduction in progeny survival, but their effects were similar across all tree densities and elevations during egg and larval life stages. However, cocoon-stage survival did vary between tree densities and elevations due to natural-enemy effects.
5. It was concluded that the observed oviposition preference for, and higher progeny performance on, trees at low densities and bottom slope elevations were caused primarily by host-plant effects.
6. These results further the argument that heterogeneity at the resource level (i.e. bottom-up forces) determines potential outcomes of multitrophic level interactions.     

Management of diamondback moth,Plutella xylostella (Lepidoptera: Plutellidae) by mating disruption

Abstract Field trials were conducted in China in 2008 and 2009 to evaluate the efficacy of mating disruption (MD) on diamondback moth, Plutella xylostella, in cabbage, Brassica oleracea var. capitata. Effectiveness was positively correlated with the MD dispenser density in the field. A density of 167 MD dispensers per ha produced an average population decrease of about 50% compared to the conventional‐practice field. Significant fewer males were captured in pheromone‐treated and conventional‐practice fields than in the blank control field, but the difference was not significant between the pheromone‐treated and conventional‐practice fields. In addition, fewer eggs and larvae were observed in pheromone‐treated fields. Our results suggest mating disruption coupled with minimal insecticidal supplements is a promising solution for resistance management and control of diamondback moth infestation.